Abstract

A study of the voltage and time dependence of a transepithelial Cl<latex>$^-$</latex> current in toad skin (Bufo bufo) by the voltage-clamp method leads to the conclusion that potential has a dual role for Cl<latex>$^-$</latex> transport. One is to control the permeability of an apical membrane Cl<latex>$^-$</latex> pathway, the other is to drive Cl<latex>$^-$</latex> ions through this pathway. Experimental analysis of the gating kinetics is rendered difficult owing to a contamination of the gated currents by cellular ion redistribution currents. To obtain insight into the effects of accumulation-depletion currents on voltage clamp currents of epithelial membranes, a mathematical model of the epithelium has been developed for computer analysis. By assuming that the apical membrane Cl<latex>$^-$</latex> permeability is governed by a single gating variable (Hodgkin-Huxley kinetics), the model predicts fairly well steady-state current-voltage curves, the time course of current activations from a closed state, and the dependence of unidirectional fluxes on potential. Other predictions of the model do not agree with experimental findings, and it is suggested that the gating kinetics are governed by rate coefficients that also depend on the holding potential. Evidence is presented that Cl<latex>$^-$</latex> transport through open channels does not obey the constant-field equation.